BENTON HARBOR POWER PLANT LIMNOLOGICAL STUDIES

PART XV: THE BIOLOGICAL SURVEY OF 12 NOVEMBER 1970

John C. Ayers
Samuel C. Mo z ley
James C. Roth

Under Contract with

American Electric Power Service Corporation
Indiana and Michigan Electric Company

Special Report No. 44
of the
Great Lakes Research Division
The University of Michigan
Ann Arbor, Michigan

July 1973

INTRODUCTION

In Part VII (March 1971) of our report series relative to the Donald
C. Cook Nuclear Station, we established the following report format:

A. COOK PLANT PREOPERATIONAL STUDIES

A.l Recording of Local Water Temperatures

A. 2 Study of Floating Algae and Bacteria

A. 3 Development of a Monitor for Phytoplankton (ABANDONED)

A. 4 Study of Attached Algae

A. 5 Study of Zooplankton

A. 6 Study of Aquatic Macrophytes

A. 7 Study of Benthic Organisms

A. 8 Study of the Local Fishes

A. 9 Support of Aerial Scanning

A. 10 Study of Entrainment and Impingement

B. SURVEYS OF EXISTING WARM WATER PLUMES

C. THE ICE BARRIER AT THE COOK PLANT SITE

D. EFFECTS OF EXISTING THERMAL DISCHARGES ON LOCAL ICE BARRIERS

E. EFFECTS OF RADIOACTIVE WASTES IN THE AQUATIC ENVIRONMENT

E.l Gamma Scan of Bottom Sediments (FINISHED)

E.2 The Most Sensitive Organism for Concentration of

Radwastes (FINISHED)
E.3 Study of Lake Michigan's Present Radioactivity Content

(FINISHED)

This report covers only items A. 2, A. 5, and A. 7 of the above format.
These studies constitute a survey of the large-scale set of biology stations
related to the Donald C. Cook Plant carried out on 12 November 1970.

The layout of sampling stations, with indication of how the stations
are numbered, is given in Figure 1. The sampling stations, their positions
relative to the Cook Plant, their distances offshore, and the water depths
encountered are given in Table 1 and Table lA.

-1-

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-2-

Table 1. The sampling stations, their positions relative to the Cook Plant,
their distances offshore, and the water depths encountered on 12 November
1970.

station

Position relative to the Cook Plant

Water Depth

(ft)

DC-1

Directly off the plant.

1/4 mi offshore

*

DC-2

3/4 "

II

45.5

DC-3

1

1/4 "

11

55.5

DC-4

It It ti 2

1/4 "

II

66.1

DC-5

.1 ti ,1 ^

It

79

DC- 6

tt ti „ 7

It

128

NDC-.25-1

1/4 mi north of the plant, 3/4

mi offshore 45

NDC-.5-1

1/2

It It II II It

1/4

II 1

16.8

NDC-.5-2

It

It It It It It

1/2

It 1

30

NDC- . 5-3

It

It 11 It It It

1 1/4

II 1

60

NDC-1-1

1

It II It It It

1/4

It 1

20

NDC-1-2

II

It It It It It

3/4

It 1

' 29.2

m)C-l-3

II

It It It It It

2 1/4

II 1

67

NDC-2-1

2

tt It It tt 11

1/4

II 1

20

NDC-2-2

II

II II II It It

1/2

It 1

' 30.2

NDC-2-3

II

It II It It II

1 1/4

It 1

49.7

NDC-2-4

II

11 It It It It

4

II 1

79.5

NDC-4-1

4

11 It It It It

1/4

It 1

18.6

NDC-4-2

II

It It It It It

1/2

ti 1

30.7

NDC-4-3

It

It It 11 ti It

2 1/4

11 1

60.5

*Station not occupied. Dredge working there.

-3-

Table 1 continued

Station

Postion relative to the Cook Plant

NDC-4-4

4

mi north

NDC-7-1

7

II II

NDC-7-2

11

II II

NDC-7-3

ft

II II

NDC-7-4

tf

It II

NDC-7-5

?t

II II

SDC-.25-1

1/4

" south

SDC- . 5-1

1/2

II II

SDC-.5-2

II

II It

SDC-.5-3

II

II II

SDC-1-1

1

It It

SDC-1-2

II

It It

SDC-1-3

II

It It

SDC-2-1

2

It If

SDC-2-2

II

It It

SDC-2-3

It

It It

SDC-2-4

"

It It

SDC-4-1

4

n It

SDC-4-2

II

It It

SDC-4-3

II

II It

SDC-4-4

II

It It

SDC-7-1 -

7

It It

It It

mi north of the plant, 7 mi offshore
It It -^/^ It It

It It -1/2 i» "

1 1/4 "

2 1/4 "
4

I 11 II ^ji^ II

, ,1 1, 3^/4 II

I It It -I I J It

1 1/4 "
I II II -^ji^ II

' " " 3/4 "

2 1/4 "
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I II II 1 yo "

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I II It 1/2 "

It It It 2 1/4 "

It If It -7

If ft ft ]^/4 II

II 11

Water

depth

(f

t)

139

19

27

46.

5

54

76

44

19

32.

3

52

25.

,3

31.

,8

64

14.

.8

30.

.4

53.

.5

74,

.6

17.

.3

29

.7

59

.5

106

.5

20

.2

-4-

Table 1 continued

Station

Position vi

ilative

to the Cook Plant

Water Depth
(ft)

mi south of

plant ,

1/2 mi offshore

30

II 1? II

II

1 1/4 "

50.6

II n II

II

2 1/4 "

55.1

II II 11

II

4

70.5

SDC-7-2
SDC-7-3
SDG-7-4
SDC-7-5

Table lA. Additional stations for phytoplankton only. (all in 4 ft.
of water.)

Station

Position relative to the Cook Plant

NDC-.5-0

1/2 mi north of the plant, just off the beach

NDC-1-0

1

II II II II II II It II 11

NDC-2-0

2

II II II II II II II II II

NDC-4-0

4

II II II II II II II II 11

SDC-.5-0

1/2

" south " " " " " " "

SDC-1-0

1

II II II II II It 11 II II

SDC-2-0

2

II II II II II II It It II

SDC-4-0

4

II II It It II It It It II

-5-

Phytoplankton samples were taken at all the stations of Table 1. At all
stations with serial numbers greater than zero, zooplankton, benthos, and
physical measurements were collected as well. The physical measurements con-
sisted of surface water temperature, water depth, bottom types, Secchi disc
water transparency, and water color as seen above the white 20-cm Secchi
disc. Weather conditions and wind and wave characteristics were taken, and
meteorological data taken on 12 November 1970 apply to all the sections of
this report; these data are presented in Appendix A.

-6-

A. 2 Study of Floating Algae and Bacteria

On the date of this survey, techniques for the determination of bacteria
were being tried but usable results were not being obtained.

The phytoplankton collections of 12 November 1970 were collected and
preserved with Utermohl's iodine fixative in our usual manner, but for
reasons not known the collections disintegrated and were unusable.

-7-

A. 5 Study of Zooplankton

Zooplankton Techniques

Zooplankton collections were made by a vertical haul, from bottom to
surface, with a #5-mesh (0.282-inm average openings) net of .5-m diameter.
A propeller-type flowmeter was affixed in the center of the net mouth to
obtain quantitative measurement of the volume of water sampled by the net.
The volume of water that passed through the net was indicated by the number
of revolutions made by the flowmeter propeller; this figure was recorded
and later converted to an equivalent expressed in liters of water.

The net was then raised above the surface and rinsed to free organisms
impacted on the net and to concentrate the sample in the collecting jar tied
on the narrow cod-end of the net. Then, excess water in the brimful jar was
decanted through a small area of the net just above the cod-end. This small
area of the net was then rinsed carefully to wash all zooplankters into the
collecting jar with a minimum amount of water. The jar was removed from
the net, and Koechies fixative, a solution of formalin and sugar, was added
as a preservative. An identification label containing pertinent collection
data was placed in the jar. The jar was capped and labeled exteriorly for
delivery to the laboratory.

In the laboratory, the sample volume was measured by transferring the
entire sample to a graduated cylinder. The entire sample then was returned
to the collecting jar and mixed thoroughly and continuously with a magnetic
stirrer while 1-ml subsamples were extracted with a Henson-Stempel pipette.
Each subsample was placed in a depression in a clear glass spot plate. Each
depression received a few drops of soap solution to break the surface tension

-8-

and allow the zooplankton to settle to the bottom for easier counting. A
variable-magnification binocular microscope was used with transmitted light
for counting and identification. As many 1-ml subsamples were counted as
were necessary to obtain good statistical parameters. The number of zoo-
plankton per liter of water was obtained by conversion with standard factors.

The #5 plankton net does not quantitatively collect the smaller crusta-
cean species, and its use was abandoned after 1970; a #10 mesh net has been
used since. The 1970 samples were enumerated by slightly different methods
than those now in use, but both methods give comparable results. Samples
counted earlier and recently recounted (given in Table 36 of Part XIII and
repeated here as Table 2) agreed closely.

The qualitative composition of the November 1970 plankton fauna is
illustrated by the species counts for stations DC-2 and DC-6 (Table 3) .
These were counted recently by current methods.

The November 1970 fauna resembled closely that reported for 1972. The
absence of nauplii and Tropocyalops prasinus from these stations can be
explained by the coarse mesh net used in 1970. We found no specimens of
Cyclops vernaliSj Chydorus sphaerious^ or Polyphemus peoiaulus^ but these
species were rare or absent in November 1972 as well. In 1970, as in 1972,
immature copepods were major components of the zooplankton assemblage (the
1970 counts are probably underestimates because of the loss of early instars
through the net). But the two bosminid species, Bosmina longirostris and
Eubosmina coregoni were more abundant in 1970, especially at the inshore
station; evidently the autumn bosminid pulse lasted longer in 1970 than in
1972. In both years Eubosmina outnumbered Bosmina in November. In 1970,
bosminids were the dominant zooplankters at all but a few offshore stations

-9-

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-10-

Table 3. Zooplankton species counts (ind/m ), coefficients of variation
between duplicate subsamples, percent composition by species, total
zooplankton weight (mg/in3), and mean zooplankter weight (yg/ind) for
2 stations sampled on 12 November 1970.

SPECIES

DC-2

#/m-

c.v.

#/m-

DC-6

c.v.

2,907
1,147

Copepod nauplii
Cyclopoid copepods
Immature copepodids

Cyalops biauspidatus thomasi

Cyclops vemalis

TTopooyotops prasinus mexioanus
Calanoid copepods

Immature copepodids

Diaptomus ashlandi

Diaptomus minutus

Diaptomus oregonensis

Diaptomus sioitis

Episahura laaustris

Eurytemova af finis

Limnoaalanus macrurus
Harpacticoid copepods

Canthooamptus sp.
Cladocerans

Bosmina longirostris

Ceriodaphnia quadrangula

Chydorus sphaeviaus

Daphnia galeata mendotae

Daphnia vetroaurva

Diaphanosoma leuohtenbergianum

Eubosmina ooregoni

Hotopedium gibberum

Leptodova kindtii

Polyphemus pediaulus
Rotifers

Asplanohna sp.

TOTAL 23,520

2,120
13

307
453

10,013
80

27

4
3

12.4
4.9

1,390
661

20
144

6
8

8

9.0
0.1

399

1.3
1.9

42.6
0.3

275

372

28

1,252

41

14

0.1 55
12,731

10

14
16

23
47
143

10.9
5.2

4,013

5

17.1

4,

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3

36.3

227

25

1.0

785

27

6.2

640

6

2.7

1,

,335

16

10.5

1,347

7

5.7

1,

,321

3

10.4

13

144

0.1

28

0.2

13

144

0.1

41

47

0.3

187

20

0.8

69
41

28
47

0.5
0.3

3.1

2.2
2.9
0.2
9.8
0.3
0.1

0.4

-11-

where they were outnumbered by immature copepods. However, the percent
composition estimates are based on totals which do not include the smaller
forms included in subsequent surveys.

No gravid copepods were noted in these samples. Several of the
cladoceran species gave evidence of fall breeding, however. Males of Daphnia
gdleata mendotae^ D. retroourva^ Diaphanosoma leuohtenbergianum^ Eolopedvum
gibberim^ and Eubosmvna oovegorvl were noted, as were ephippial females of
Daphnia retroaurva and both bosminid species.

The counts for the other stations (enumerated by the earlier method)
are given in Table 4. We have also listed the diversity indicies calculated
for each station; these are included for comparative purposes since this
parameter was reported for earlier collections. These calculations are
subject to the criticism that they are based on superspecif ic categories
collected with size-selective apparatus. Other objections are possible as
well. Recent authors (Hurlbert 1971; Hill 1973) have criticized the concept
of the diversity index and questioned its usefulness as an ecological in-
dicator. And the supposed relationship between "diversity" and "stability"
in ecosystems remains to be demonstrated.

Total zooplankton numbers collected in November 1970 were roughly
comparable to those found in 1972. A few stations near the plant site and
south of it (circled in Figure 2) had abundances of over 30 individuals per
liter. The spatial distributions of the three most abundant zooplankton
groups on 12 November 1970 are summarized in Figures 2-5. Bosminids were
concentrated near shore (Figure 3) , where they usually numbered between 8 and
20 per liter; two inshore stations had over 50 per liter, and four had 5-6
per liter. All stations offshore of the heavy line in Figure 3 had less

-12-

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-16-

Table 4. Zooplankton, 12 November 1970. Samples by vertical haul of
metered #5 Net. Organisms per liter.

Organisms

*

DC-1

DC-2

Stat
DC-3

ions
DC-4

DC-5

DC-6

Copepods:

Diaptomids

—

5.34

6.05

3.31

4.61

8.81

Epischura

—

0.11

0.12

0.04

0.04

0.05

Eurytemora

—

—

0.08

—

—

0.02

Limnocalanus

—

—

—

—

—

0.02

Senecella

—

0.02

—

0.03

—

—

Cyclopoids

—

4.57

8.73

3.09

4.04

2.18

Harpactacoids

—

0.02

—

—

—

—

Cladocerans:

Alona

—

—

—

—

—

—

Bosmina

—

14.99

15.22

10.06

9.13

2.00

Ceriodaphnia

—

—

—

0.01

—

—

Daphnia

—

0.97

1.12

0.79

0.59-

0.64

Diaphanosoma

—

0.07

0.11

—

—

0.01

Eurycercus

—

—

—

—

—

—

Holopedium

—

0.15

0.21

0.17

0.08

0.03

Leptodera

—

0.04

0.03

0.01

0.03

—

Polyphemus

—

0.04

—

—

—

—

Rotifers:

Asplanchna

0.04

0.08

0.04

0.03

0.01

TOTAL

26.36

31.75

17.55

18.55

13.77

Diversity index

1.70

1.81

1.69

1.72

1.53

*Station not occupied, dredges on the position.

-17-

Table 4 continued

Stations

Organisms

NDC-.25-1

NDC-.5-1

NDC

:-.5-2

NDC-.5-3

NDC-1-1

Copepods:

Diaptomids

2,40

1.76

*

*

2.27

Epischura

0.06

0.04

*

*

0.06

Eurytemora

0.11

0.02

*

*

—

Limnocalanus

0.04

—

*

*

—

Senecella

0.04

—

*

*

—

Cyclopoids

1.84

3.06

*

*

8.35

Harpactacoids

—

—

*

*

—

Cladocerans:

Alona

—

—

*

*

—

Bosmina

6.20

10.06

*

*

52.71

Ceriodaphnia

—

—

*

*

—

Daphnia

0.44

0.65

*

*

1.23

Diaphanosoma

0.02

0.02

*

*

—

Eurycercus

0.02

—

*

*

—

Holopedium

0.06

0.10

*

ic

0.32

Leptodera

—

0.01

*

•k

—

Polyphemus

—

—

*

-k

—

Rotifers:

Asplanchna

0.01

0.02

*

•k

0.26

TOTAL 11.24 15.74 65.20

Diversity index 1.80 1.53 0.98

*Sample broken

-18-

Table 4 continued

Stations

Organisms

NDC-1-2

NDC-1-3

NDC-2-1

NDC-2-2

NDC-2-3

NDC-2-4

Copepods:

Diaptomids

2.98

3.86

3.61

4.85

3.23

7.66

Epischura

0.03

0.04

0.03

—

0.06

0.10

Eurytemora

0.02

0.11

—

0.03

0.14

—

Limnocalanus

—

0.01

—

—

—

—

Senecella

—

0.10

—

—

0.02

—

Cyclopoids

4.36

4.82

3.81

4.45

6.18

3.08

Harpactacoids

—

0.01

—

——

Cladocerans :

Alona

—

0.01

—

Bosmina

16.13

9.09

19.09

11.81

8.38

4.12

Ceriodaphnia

—

—

—

—

0.02

—

Daphnia

0.66

1.20

0.71

0.67

0.50

0.71

Diaphanosoma

0.07

—

—

0.03

0.05

—

Eurycercus

—

—

—

—

—

—

Holopedium

0.17

0.20

0.28

0.13

0.11

0.06

Leptodera

0.02

0.02

—

—

—

—

Polyphemus

—

0.01

—

——

"-""

"~"~

Rotifers:

Asp lane hna

0.19

0.03

0.06

0.03

0.05

0.04

TOTAL

24.63

19.51

27.59

22.00

18.74

15.77

Diversity index

1.51

1.96

1.38

1.67

1.81

1.77

-19-

Table 4 continued

Stations

Organisms

NDC-4-1

NDC-4-2

NDC-4-3

NDC-4-4

NDC-7-1

NDC-7-2

Copepods:

Diaptomids

1.21

1.55

5.78

9.63

5.24

3.54

Eplschura

0.04

—

0.05

0.07

0.03

0.05

Eurytemora

0.04

—

0.01

0.01

—

0.04

Limnocalanus

—

—

—

0.01

—

—

Senecella

—

0.02

—

—

—

—

Cyclopolds

4.29

3.32

2.41

1.99

6.89

4.03

Harpactacoids

—

—

—

—

—

Cladocerans :

Alona

—

—

—

—

—

—

Bosmina

21.62

10.75

2.15

2.25

12.13

11.04

Cerlodaphnia

0.04

0.02

—

—

—

—

Daphnia

0.33

0.64

0.39

0.69

0.75

0.48

Diaphanosoma

0.25

—

—

0.02

—

—

Eurycercus

—

—

—

—

—

—

Holopedium

0.29

0.11

—

0.08

0.38

0.17

Leptodera

—

0.04

—

—

0.03

0.04

Polyphemus

0.04

0.04

—

—

0.03

0.05

Rotifers:

Asplanchna

0.08

0.04

0.01

0.04

0.07

0.01

TOTAL

28.23

16.53

10.80

14.7 9

25.55

19.45

Diversity Index

1.18

1.51

1.66

1.54

1.79

1.66

-20-

Table 4 continued

Stations

Organisms

NDC-7-3

NDC-7-4

NDC-7-5

SDC-.25-1

SDC-.5-1

SDC-.25-2

Copepods:

Diaptomids

4.92

5.71

9.75

4.82

3.10

*

Epischura

0.07

0.04

0.08

0.05

—

*

Eurytemora

—

0.03

0.03

*

—

*

Limnocalanus

—

—

—

*

—

*

Senecella

—

—

—

*

—

*

Cyclopoids

3.60

2.61

2.40

3.93

5.30

*

Harpactacoids

—

—

0.01

—

*

Cladocerans :

Alona

—

—

—

—

*

Bosmina

9.90

1.88

3.18

11.00

25.90

*

Ceriodaphnia

—

—

—

—

—

*

Daphnia

0.46

0.20

0.73

0.61

0.88

*

Diaphanosoma

—

0.02

—

0.02

0.03

*

Eurycercus

—

—

—

—

—

*

Holopedium

0.13

0.06

0.11

0.14

0.13

*

Leptodera

0.02

—

—

—

0.03

*

Polyphemus

—

—

—

—

—

*

Rotifers:

Asplanchna

—

—

0.06

0.05

0.24

*

TOTAL

19.10

10.55

16.35

20.62

35.61

Diversity index

1.67

1.64

1.65

1.68

1.28

*Sample broken

-21-

Table 4 continued

Stations

Organisms

SDC--5-3

SDC-1-1

SDC-1-2

SDC-1-3

SDC-2-1

SDC-2-2

Copepods:
Diaptomids
Epischura

*
*

3.58

2.89
0.02

7.09
0.10

1.88
0.08

4.10
0.04

Eurytemora

*

—

—

—

0.01

Limnocalanus

*

—

—

0.03

Senecella

*

—

—

—

Cyclopoids

*

13.80

4.55

5.44

2.48

2.96

Harpactacoids

*

'

"

Cladocerans :

Alona

*

—

—

—

—

Bosmina

*

14.50

18.73

23.28

13.48

9.21

Ceriodaphnia

Daphnia

Diaphanosoma

*

0.91

1.07
0.02

1.21
0.03

0.96

0.70
0.03

Eurycercus
Holopedium
Leptodera
Polyphemus

*
*
*
*

0.37
0.05

0.47
0.02
0.02

0.28
0.03

0.60
0.04

0.26
0.03

Rotifers:

Asplanchna

*

0.05

0.12

0.01

—

TOTAL

33.26

27.91

37.50

19.52

17.34

Diversity index

1.64

1.49

1.55

1.49

1.75

*Sample broken

-22-

Table 4 continued

Stations

Organisms

SDC-2-3

SDC-2-4

SDC-4-1

SDC-4-2

SDC-4-3

SDC-4-4

Copepods:

Diaptomids

26.29

19.21

3.42

9.33

11.26

8.58

Epischura

0.26

0.31

0.03

0.10

—

0.05

Eurytemora

0.11

0.03

—

0.05

—

—

Limnocalanus

—

—

—

—

—

0.02

Senecella

—

—

—

—

0.15

0.02

Cyclopoids

16.99

14.89

2.13

8.07

8.07

4.58

Harpactacoids

—

— —

— —

"•"■

Cladocerans :

Alona

—

—

Bosmina

59.29

17.54

5.41

10.67

14.73

4.10

Ceriodaphnia

—

—

—

—

—

—

Daphnia

1.95

2.89

0.56

1.58

1.14

0.83

Diaphanosoma

0.11

—

—

0.10

—

0.05

Eurycercus

—

—

—

—

—

—

Holopedium

0.79

0.31

0.07

0.37

0.31

0.11

Leptodera

0.15

—

—

—

—

—

Polyphemus

— —

—

—

—

0.02

Rotifers:

Asplanchna

0.15

0.14

0.17

0.05

0.09

0.12

TOTAL

106.09

55.32

11.79

30.32

35.75

18.48

Diversity index

1.62

1.90

0.88

1.95

1.81

1.87

-23-

Table 4 continued

Organisms

SDC-7-1

SDC-7-2

Station^
SDC-7-3

SDC-7-4

SDC-7-5

Copepods:

Diaptomids

8.05

2.50

4.63

13.89

8.17

Epischura

0.17

0.01

0.08

0.24

0.05

Eurytemora

—

0.02

0.03

0.05

—

Limnocalanus

—

—

—

—

—

Senecella

—

—

—

—

—

Cyclopoids

7.31

1.81

2.52

5.97

7.95

Harpactacoids

—

—

—

—

—

Cladocerans :

Alona

—

—

—

—

—

Bosmina

13.47

5.55

4.67

10.02

3.05

Ceriodaphnia

—

—

—

—

—

Daphnia

0.96

0.20

0.64

0.91

0.72

Diaphanosoma

—

0.02

0.03

—

—

Eurycercus

—

—

—

—

—

Holopedium

0.20

0.11

0.07

0.19

0.09

Leptodera

0.03

—

—

—

0.04

Polyphemus

—

—

—

—

—

Rotifers:

Asplanchna

0.03

0.03

0.03

—

—

TOTAL

30.22

10.25

12.70

31.27

20.07

Diversity index

1.79

1.67

1.89

1.76

1.72

-24-

than 5 individuals per liter. Diaptomids in general were most abundant
offshore (Figure 4); all stations offshore of the heavy line in Figure 4
had 7 or more individuals per liter, whereas all stations shoreward of the
line had 1-6 per liter. Highest diaptomid abundances (11-26 per liter)
occurred at four of the stations south of the plant site (enclosed by the
broken line in Figure 4) which had the highest total zooplankton abundances.
Cyclopoids occurred throughout the study area in concentrations between 2
and 5 individuals per liter (Figure 5) , except for a few scattered nearshore
stations and six stations south of the plant site (circled in Figure 5) ,
where 6-17 per liter were found. These distributions suggest a large patch
of plankton-rich (offshore?) water occurred south of the plant site on
12 November 1970.

-25-

A. 7 Study of Benthic Organisms

Benthos Techniques

Benthic organisms were collected by use of the Ponar grab-sampler.
Two grabs were combined and passed together through a washing device in
which the benthic organisms were retained on a 0.5-mm mesh screen. In sub-
sequent counting, the counts were divided by two to give the average of the
duplicate samples. Organisms from the washing device then were collected
into pint Mason jars, labeled internally and externally, preserved with
buffered formalin, and returned to the laboratory for processing. In the
laboratory, the samples were concentrated on a small mesh net, and trans-
ferred with minimum fluid to the counting tray.

For general survey purposes, the benthos are counted into the groups:
amphipods, oligochaetes, sphaeriids, chironomids, and others (mostly leeches
and snails). The averaged counts were converted by standard factors to give
numbers of organisms per square meter. The counted samples are preserved by
appropriate standard museum techniques and retained as a reference collection.
Initially, another compromise was needed to expedite enumeration of the
oligochaetes. These worms tend to fragment during processing, and it was
not possible at first for us to rapidly distinguish fragments from whole
individuals. Therefore, to estimate oligochaete abundance, all worms and
parts of worms were counted, and the total divided by three. More detailed
examination of samples has shown that this factor varies from sample to
sample with the result that oligochaete abundances obtained by this method
are fairly good estimates rather than real counts. Oligochaetes are now
counted by head ends, and ignoring other fragments (literally counting heads).

-26-

Inconsistencies between abundances given in Table 5 and in Table 39 of Part
XIII are due largely to head counts being used in this report while frag-
ments/3 was used in Part XIII Tables 38 through 43. All other tables in
Part XIII are based on the head count method. Table 5 gives, by depth,
the percent of population comprised by major taxa and also the total numbers
of organisms per square meter as well as the numbers of taxa identified and
the diversity indices computed from species counts for each of the 35 stations
sampled.

This report details the species compositions (Table 6) and other as-
pects of 35 benthos samples from stations in all parts of the survey area,
all based on the oligochaete head count method.

Benthos Abundances

Many aspects of the November 1970 benthos data were reported in Part
XIII of this report series and are here repeated for the reader's conven-
ience: average abundances and frequencies of occurrence of species for
all stations combined (Part XIII, Table 47) is here Table 7; percentage
composition by depth zones (XIII, Table 49) is here Table 8; and average
abundances of dominant taxa by benthic (depth) zones (XIII, Table 51) is
here Table 9. Other tables and figures here included are specific unto
this report.

In comparison to July 1970, the November 1970 collections showed greater
abundances of Tubif icidae and Pisidium^ while Pontoporeia affinis was less
abundant (Table 7). The pattern of depth distribution and benthic zonation
was, however, little affected by the shifts in dominant taxa abundances
that took place between July and November (Tables 8 and 9) .

-27-

Species div-
ersity index

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-46-

Table 8. Contribution of dominant taxa to the benthos community.

% of Total Fauna
Jul 1970 Nov 1970 Apr 1971 Jul 1971

Pontoporeia affinis

49.9

22.8

33.8

37.6

Stylodritus heringianus

14.8

6.7

22.1

11.5

Tublficidae

16.3

43.3

24.2

33.8

Sphaerium nitidum

4.2

2.4

2.9

2.0

P-tsidium spp.

8.6

17.2

12.2

9.0

Chironomidae

4.7

6.8

3.6

5.2

Remainder

1.5

0.8

1.2

0.9

Total #/m

3528

5906

3641

6810

-47-

Table 9. Mean numbers by depth zone, for species dominant at some
time of year, on 12 November 1970.

Species 0-8m 8-16m 16-24m >24m

Pontoporeia affinis
Stylodrilus heringianus
Limnodrilus hofftneisteri (mature)
Limnodrilus angustipenis (mature)
PetosQolex freyi (mature)
Sphaeriim nitidim
Pisidium spp.

Chironomus anthracinus-gr.
Cryptochironomus sp. 2
Paraaladopelma nereis
Paraohironomus cf. demeijerei
Polypeditum cf. soalaenum

15

214

1,584

5,345

68

360

1,664

14

725

164

93

78

53

35

36

384

36

100

985

1,105

1,977

140

17

4

36

111

3

5

1

-48-

Figure 6 shows the total numbers of benthic macroinvertebrates, plotted
against depth, for the 35 selected samples. Stations near shore are coded
for their distance from shore. It is clear that stations near shore had very
few animals. Proceeding away from shore, the average number of individuals
increased, but the variability among stations at similar depths increased
also. The few stations over 30 meters deep seemed to have somewhat more
uniformly high abundances. There was no consistent tendency for stations in
the center of the survey area to have more benthos than stations farther north
or south. This was different from July, when stations in front of the plant
had more benthos than stations farther north and south.

Benthic Zone (0 to 8 m)

All of these stations were 1/4 mile (400 m) from shore, and together

2
averaged 332 organisms per m . Tubificidae (Limnodrilus hoffmeisteri) with

38% of the population, Pisidium with 30%, and Chironomidae with 27% were the

major component forms. Chironomus fluviatilis-group and Cryptoohironomus spp.

made up most of the Chironomidae. Two species which were abundant or frequent

in the 10 July 1970 collections from this zone, Paraaladopelma nereis and

Paraohironomus cf. demeijerei (Mozley and Garcia 1972), were nearly absent in

November. These midges had emerged from the lake in summer and the next

generation had not yet grown to a size which could be retained by our 0.5r-mm

sieve in November.

Benthic Zone 1 (8 to 16 m)

This benthic zone, including mostly stations at 1/2 to 3/4 mile (800 and

1200 m) from shore, had many more animals. The average total abundance was

2
2,556 organisms per m . One sample, SDC-7-3, was anomalous (see below) and

-49-

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-50-

was excluded from this average and from following percentages. The major
taxa in this zone were still more heavily dominated by oligochaetes (48%) ,
and several additional species were present in abundance: Potamothrix
moldaviensiSj other Limnodrilus species, Aulodrilus amerioanus^ Potamothria:
vejdovskyij and (in the deeper stations of this zone) Stylodrilus heringianus
and immature Tubificidae with hair setae (mainly Tubifex tubifex^ but possibly
including Ilyodrilus templetoni) . The Chironomidae (16% of the population)
were again dominated by Chironomus and CryptoahironomuSj but several other
species contributed to the total in the deeper part of this zone: Chironomus
anthracinus-group, Prooladius^ Kiefferulus^ Monodiamesay and Potthastia.
Polypedilum cf . saalaenum was common in this zone on 10 July 1970 but must
have emerged from the lake in the intervening months, Pisidivm^ at 21% of
the population, comprised a smaller percentage but was more abundant in
absolute terms than near the beach. Several species of Sphaerium were found.
Pontoporeia averaged 9% of the population and was present in all samples from
this zone; it was more abundant here than in the beach zone, but was still
only a minor part of the benthos.

Benthic Zone 2 (16 to 24 m)

The third benthic zone, composed mostly of stations between 1-1/4 to
2-1/2 miles (2-4 km) from shore, was not very different in benthos species
from the zone described above except that the numerical fraction due to
Pontoporeia increased to an average of 26%; Sphaerium nitidum at 6% of the
population and Stylodrilus at 6% of the population also represented increases.
The rises in numbers, primarily among these three animals, produced an
increase in average total abundance to 5,993 organisms per square meter.
Limnodrilus hoffmeisterij Limnodrilus cervix ^ Aulodrilus spp., Chironomus spp..

-51-

and Cryptochironomus sp. 2 decreased in this zone relative to benthic zone 1.

Benthic Zone 3 (more than 24 m)

The deepest part of the survey area was characterized by relative increases
±n Pontoporeia (to 54% of the population) and of Stylodrilus (to 16%) and by
declining numbers of other species. The average total abundance of benthos
was 10,362 organisms per square meter.

Station SDC-7-3

One station (SDC-7-3) stood out from the rest because of its sample size
and species composition. This station produced extremely large populations
of several species of pollution-tolerant Tubificidae, and many Prooladius and
Fisidivim. Frooladius is believed to be a predator on oligochaetes, and is
tolerant of organic pollution. The number of total oligochaetes and the large
proportion of mature Limnodrilus cervix were similar to some samples from the
Toledo area of western Lake Erie and far exceeded the abundances of Oligochaeta
previously reported from Lake Michigan. This station is not near any major
source of pollution, and it is a mystery how such a benthic association could
develop. It is considered too anomalous to include in overall averages of
benthos in the survey area.

Species Diversity of Benthos

Species diversity indices which combine richness in species with evenness
of distribution of individuals among species have recently become established
as important tools in the definition of polluted areas in streams (Wilhm and
Dorris 1968) . Their use is now being extended to preoperational surveys in
Lake Michigan (Mozley and Garcia 1972; Beak Consultants, Inc. 1973). If this
is to be done rationally, the factors which influence diversity in Lake Michigan

-52-

must be considered in some detail. On the basis of November 1970 benthos
data, we contend that the interpretation of species diversity measurements
will be difficult because of the many biological and environmental factors
which influence them.

Species diversities were calculated for the 35 selected samples from
the November 1970 survey which were identified to species (Table 5) . The
formula for species diversity:

d = -Z (N^/n log^ N^/n)
is derived from Shannon and Weaver (1963) . Certain conventions were adopted
for the calculation of this index. First, the genus Pisidium was counted as
a single species as it has not yet been sorted into species. Since it is
often one of the most numerous taxa, splitting it into species would increase
the diversity indices somewhat. Several Pisidium species are abundant.
Secondly, many Tubificidae cannot be identified to species before they are
mature. For calculation of diversity indices we have divided the immatures
in each sample or combination of samples in the same proportion as mature
Tubificidae to which they could belong. Some species of Oligochaeta, such as
Aulodrilus spp., Potamotkrix vejdovskyij Pelosoolex multisetosus^ and
Stylodrilus heringianus^ can be identified in all stages after the egg, so
they were excluded from the apportionment of immatures.

The diversity indices were slightly higher in November 1970 than in July
1970 (Table 10), except near the beach in benthic zone 0. In this zone the
chironomids abundant in July were rare or absent in November. In the deeper
zones Pontoporeia was less abundant and a mixture of Tubificidae and Pisidium
were more abundant, making the distribution of individuals among different
taxa more uniform. The range of diversity indices was again very wide.

-53-

Table 10. Ranges and averages of benthos species diversity in the Cook Plant
survey area in Julv and November 1970.

Month

Overall
average

Maximum

Minimum

Averages by benthic zones
Zone Zone 1 Zone 2 Zone 3

July
November

2.17
2.26

3.4
3.34

0.9
0.65

1.96
1.52

2.54 2.23 1.40
2.64 2.43 1.78

including values which in streams would indicate severe pollution (less than
1.0) or very clean water (more than 3.0). Highest diversities, on the average,
were in benthic zone 1, which was the richest in taxa. It had 32 total taxa,
and an average of 13.6 taxa, compared to 14.1 average but only 27 total taxa
in the next most diverse benthic zone, 2. Benthic zone 3 had a total of only
14 taxa, but an average of 9.8 taxa per sample, which illustrates the increase
in homogeneity of the fauna in the profundal zone of Lake Michigan. The fewest
taxa were in benthic zone 0, which had a total of 10 distinguishable taxa, and
an average of 4.2 taxa per sample.

The difference between zones is also related to the dominance of Tontoporeia.
In benthic zone 1, Pontoporeia accounted for only 3.2% of the combined total
abundances. Excluding the atypical sample (SDC-7-3) described in the preceding
section, Pontoporeia made up 8.8% of the total abundance. In zone 2, however,
Pontoporeia comprised 26% of the total abundance on the average, but was much
higher in some samples (Table 6). The diversities in the latter zone conse-
quently tended to be lower. The effect of numerical dominance by Pontoporeia
on species diversity is illustrated in Figure 7. As its percentage of the
total population goes above 30%, the diversity index decreases. The reduction

-54-

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-55-

in the index due to the small nxomber of taxa which tolerated the instability
of the beach zone is also illustrated there.

Johnson and Brinkhurst (1971) recognized the same effects on species
diversity of Lake Ontario benthos, and attributed them to cold water and
reduced environmental heterogeneity. A protected warmer embayment, even
where it was organically enriched, had higher indices of diversity than the
open lake.

In the Great Lakes man's activities could modify species diversity. For
instance, unspecified types of pollution are presumed to have reduced the
abundance (or at least the dominance) of Pontoporeia and increased the
abundance of oligochaetes in southern Lake Michigan (Powers and Robertson
1965; USFWPCA 1968). Since the Oligochaeta comprise many species which tend
to increase together, pollution could increase species diversity at least
initially. The additional habitats offered by intake and outfall structures,
breakwalls and other man-made devices increase the diversity of the environ-
ment and perhaps that of the population of benthos.

On another tack, the diversity index of a sample is not necessarily
representative of the local community. In samples from shallower depth zones
(see text above) , only half or less of all species present occur in an
average sample, and the proportion of taxa in different samples varies widely
(Table 5).

It could be argued that rare species, whose effects on this particular
diversity index are smiall (Wilhm and Dorris 1968), cause this discrepancy
and therefore their absence from samples should not affect diversity very
much. Table 11 gives sets of species diversity indices calculated in various
combinations from five samples from the November 1970 collections. The

-56-

Table 11. Composite and average species diversities for five stations in the
center of the November 1970 survey area. All stations in benthic zone 1 (8-16
meters). Sample codes: 1 = SDC-.5-2; 2 = SDC-.25-1; 3 = NDC-.25-1; 4 = DC-2;
5 = SDC-.5-3.

Number of

Composite

Average

samples

Samples

by

Sums of -
abundance/m

Number

species

species

combined

code

of taxa

diversity

diversity

1

1

1,962

8

2.17

2

1,116

11

2.49

3

5,531

21

3.04

4

2,908

21

3.34

5

2,908

14

2.92

2

3.5

8,439

22

3.15

2.98

5.1

4,870

15

2.88

2.55

3

2,4,5

6,932

23

3.38

2.92

2.3,5

9,555

22

3.11

2.82

A

1.2,3,5

11,517

22

3.02

2.66

1.2,3,4

11,517

26

3.16

2.76

5

1,2,3,4,

,5

14,425

27

3.28

2.79

samples all came from stations within a half mile of the central transect of
the survey area, and all were from benthic zone 1 where the average sample
diversity was highest. The samples were combined in pairs, triplets, etc.,
by selecting from the five at random using a table of random numbers. Two
different combinations were used for each number of combined samples and a
composite species diversity index was computed for each combination. For
comparison, the simple averages of diversity indices of the individual samples
are given for each combination.

The trend towards increasing diversity with larger sample sizes is not
very strong, but the composite diversity was always greater than the average

-57-

diversity. The composite diversity of all five samples combined is the best
representation of the area, but it is not very representative of the
diversity to be expected in a single sample. Since five samples in combi-
nation produced a diversity which still differed from, that of a subset of
four of them by almost 0.3 units, even four samples were insufficient to give
a precise representation of the local species diversity, and it may be that
more than five samples would be necessary. A total of 1,674 animals from
10 casts of the Ponar sampler were identified to produce the data in Table 11.
The extension of this amount of effort in only a part of the survey area to
achieve a more precise depiction of species diversity over the whole study
area is not justifiable.

A less laborious and probably equally precise summarization of species
diversity might be obtained by two other measures: the number of species
and some measure of dominance of abundant species (Hill 1973) . This would
also separate the effects of these two factors which interact in less than
obvious ways in their effects on the Shannon and Weaver index. The value
of possessing diversity information in the Great Lakes remains to be
demonstrated.

For the record, diversity indices calculated from our benthos samples
are biased toward larger species and the larger individuals of smaller
species. Many small chironomids and oligochaetes undoubtably escape through
our 0.5-mm sieve while older and larger individuals are retained and counted.
Several sorts of organisms which are present in the samples are not repre-
sented in the counts because the great majority of them pass through the
sieve (e.g. Nematoda, Harpacticoidea, Ostracoda) . Therefore, the basis for
calculation is not a complete sample of the benthic community and is of value
only in comparison to other samples collected in the same way.

-58-

The use of low species diversity indices as indicators of pollution must
be restricted to streams; it is not applicable to Lake Michigan. The per-
centages and kinds of individuals comprising the dominant species, and the
number of species in different samples are too variable. The reader's
attention is called again to Figure 7, wherein diversity indices of 1.67,
1.42, 1.20 and 0.91 were related to 56.6, 76.1, 79.1 and 83.6 percent domi-
nances of the clean-water amphipod Pontoporeia af finis ^ respectively.

The use of diversity indices which are subject to multiple influences of
environmental heterogeneity, biological instability and technological error
on an arbitrary fraction of the benthic community, in a habitat where there
is no dependable theoretical framework for interpreting index values, has
no usefulness in judging the condition of the benthic community. In Lake
Michigan samples, there may be very few taxa or an overwhelmingly dominating
taxon when the index is low; the low index value may indicate either
pollution or clean water, depending on what the composition of taxa happens
to be. The direction and magnitude of changes in benthos species diversity
indices in Lake Michigan cannot be interpreted without concomitant knowledge
of the identity of dominant taxa and what their presence indicates about the
state of the aquatic environment. There is no strong reason for continuing
to calculate this parameter from our benthos sample data.

Judgments of water quality on the basis of benthos information can be
made as well from the identity of frequently occurring species in a few
taxonomic groups as from the detailed composition of species in all taxa.
The Pericarida (Amphipoda and Isopoda) , Tubificidae, Chironomidae and a few
other aquatic insects are the only groups which have accepted usefulnees for
such judgments in the Great Lakes. First priority, therefore, should be

-59-

placed on obtaining detailed knowledge of these groups, leaving such problems
in species diversity calculation as apportionment of immature Tubificidae,
identity of Pisidium species, and loss of small specimens in sieving to
theoretical studies of community structure.

-60-

LITERATURE CITED

Beak Consultants, Inc. 1973. Evaluation of preoperational biological con-
ditions of Lake Michigan in the vicinity of the Palisades Nuclear Plant.
Report to Consumers Power Co., Jackson, Michigan, File D-2048 unnumbered.

Federal Water Pollution Control Administration. 1968. Water quality in-
vestigations. Lake Michigan basin: Biology. FWPCA, U. S. Dept. of
the Interior, Chicago, Illinois. 41 p.

Hill, M. 0. 1973. Diversity and evenness: a unifying notation and its
consequences. Ecology 54: 427-432.

Hurlbert, S. H. 1971. The nonconcept of species diversity: a critique
and alternative parameters. Ecology 52: 577-586.

Johnson, M. B. and R. 0. Brinkhurst. 1971. Associations and species di-
versity in benthic macroinvertebrates of Bay of Quinte and Lake Ontario.
J. Fish. Res. Bd. Canada 28: 1683-1697.

Mozley, S. C. and L. C. Garcia. 1972. Benthic macrofauna in the coastal

zone of southeastern Lake Michigan. Proc. 15th Conf . Great Lakes Res.,
p. 102-116.

Powers, C. F. and A. Robertson. 1965. Some quantitative aspects of the
macrobenthos of Lake Michigan. Proc. 8th Conf. Great Lakes Res.,
p. 153-159.

Saether, 0. A. 1973. Taxonomy and ecology of three new species of Mono-
diamesa Keiffer, with keys to Nearctic and Palearctic species of the
genus (Diptera: Chironomidae) . J. Fish. Res. Bd. Canada 30: 665-679.

Shannon, C. E. and W. Weaver. 1963. The mathematical theory of communi-
cation. Univ. of Illinois Press, Urbana, Illinois. 117 p.

Wilhm, J. L. and T. C. Dorris. 1968. Biological parameters for water quality
criteria. Bioscience 18: 477-481.

-61-

Master list of Benthic Macrofauna in the collections of 12 November 1970.

Amphipoda
Oligochaeta

Sphaeriidae

Chironomidae

Pontoporeia affinis

Sty lodri lus heringianus
Limnodrilus hoffmeistevi
Limnodrilus cervix
Limnodrilus angustipenis
Lirrmodvi lus profundioo la
Potamothrix moldaviensis
Potamothrix vejdovskyi
Peloscolex freyi
Pelosoolex multisetosus
Tubifex tubifex
Aulodrilus amerioanus
Aulodrilus pluriseta
Iiranatures with hair setae
Immatures w/o hair setae

SphaeriiMn striatinum
Sphaeriian nitidum
Sphaerium transverswn
SphaeriiAm sp. 2
Sphaerium sp. 3
Pisidium spp.

Chironomus an thr acinus-group
Chironomus fluviatilis-group
Kiefferulus sp.
Crypto chironomus sp • 2
Cryptochironornus sp, 3

-62-

Chironomidae (cont . )

Paracladopelma nereis
Tolypedilum cf . soalaenvm
Tanytarsini spp.
Proctadius spp.
Potthastia cf. longimanus
Monodiamesa cf. bathyphila
Hetevotvissoctadius cf. subpitosus
Heterotrissoctadius cf. grimshawi

Gastropoda

Hirudinea

Lymnaea spp.
Valvata sp.

Helobdella stagnalis
Eelobdella elongata
Glossiphonia oomplanata
Nephelopsis obscuva

-63-

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